In a later stage the openings of the two atrial tubes gradually travel backwards, and at the same time approximate, till finally they meet and coalesce at the posterior end of the blastoderm behind the nervous disc ([fig. 13], cl). The tubes themselves at the same time become slightly constricted not far from their hinder extremities, and so divided into a posterior region nearly coterminous with the nervous system ([fig. 13]), and an anterior region. These two regions have very different histories in the subsequent development.
The nervous disc has during these changes become marked by a median furrow ([fig. 13], ng), which is soon converted into a canal by the same process as in the simple Ascidians. The closure of the groove commences posteriorly and travels forwards. These processes are clearly of the same nature as those which take place in Chordata generally in the formation of the central nervous system.
In the region of the germinal disc which contains the anterior part of the atrial tubes, the alimentary cavity becomes, by the growth of the layer of cells described in the last stage, a complete canal, on the outer wall of which the endostyle is formed as a median fold. The whole anterior part of the blastoderm becomes at the same time gradually constricted off from the yolk.
Fig. 13. Blastoderm of Pyrosoma shortly before its division into Cyathozooid and Ascidiozooids. (After Kowalevsky.)
cl. cloacal (atrial) opening; en. endostyle; at. atrial cavity; ng. nervous groove.
The heart and pericardial cavity are seen to the left.
The fate of the anterior and posterior parts of the blastoderm is very different. The anterior part becomes segmented into four zooids or individuals, called by Huxley Ascidiozooids, which give rise to a fresh colony of Pyrosoma. The posterior part forms a rudimentary zooid, called by Huxley Cyathozooid, which eventually atrophies. These five zooids are formed by a process of embryonic fission. This fission commences by the appearance of four transverse constrictions in the anterior part of the blastoderm; by which the whole blastoderm becomes imperfectly divided into five regions, [fig. 14] A.
The hindermost constriction (uppermost in my figure) lies just in front of the pericardial cavity; and separates the Cyathozooid from the four ascidiozooids. The three other constrictions mark off the four Ascidiozooids. The Cyathozooid remains for its whole length attached to the blastoderm, which has now nearly enveloped the yolk. It contains the whole of the nervous system (ng), which is covered behind by the opening of the atrial tubes (cl). The alimentary tract in the Cyathozooid forms a tube with very delicate walls. The pericardial cavity is completely contained within the Cyathozooid, and the heart itself (ht) has become formed by an involution of the walls of the cavity.
The Ascidiozooids are now completely separated from the yolk. They have individually the same structure as the undivided rudiment from which they originated; so that the organs they possess are simply two atrial tubes, an alimentary tract with an endostyle, and undifferentiated mesoblast cells.
In the following stages the Ascidiozooids grow with great rapidity. They soon cease to lie in a straight line, and eventually form a ring round the Cyathozooid and attached yolk sack.